Saussurea involucrata PIP2;4 improves growth and drought tolerance in Nicotiana tabacum by increasing stomatal density and sensitivity

Aquaporins, the major facilitators of water transport across membranes, are involved in growth and development and adaptation to drought stress in plants. In this study, a plasma membrane intrinsic protein (SiPIP2;4) was cloned from Saussurea involucrata, a cold-tolerant hardy herb. The expression of SiPIP2;4 increased the stomatal density and sensitivity of tobacco (Nicotiana tabacum), thus, affecting the plant's growth and resistance to the diverse water environment. The higher stomatal density under well-watered conditions effectively promoted the photosynthetic rate, which led to the rapid growth of transgenic lines. The stomata in the transgenic lines responded more sensitively to the vapor pressure deficit than the wild-type under different levels of ambient humidity. Their stomatal apertures positively correlated with the ambient humidity. Under drought conditions, the overexpression of SiPIP2;4 promoted rapid stomatal closure, reduced water dissipation, and enhanced drought tolerance. These results indicate that SiPIP2;4 regulates the density and sensitivity of plant stomata, thus, playing an important role in balancing plant growth and stress tolerance. This suggests that SiPIP2;4 has the potential to serve as a genetic resource for crop improvement.